Penaeid and carid community changes in the St Lucia estuarine lake system,South Africa,under low water level,extended closed periods and marine reconnection conditions

Penaeid and carid communities were assessed in St Lucia over a wide range of saline conditions, water level conditions and mouth states, including a six-month marine connection and various connections to the Mfolozi Estuary. Samples were collected biannually in spring and autumn from November 2004 to May 2012 using seine nets. Seven penaeid and seven carid species were recorded, of which Palaemon pacificus, P. peringueyi, Penaeus indicus and Metapenaeus monoceros were the most abundant. The decline from 2004 to 2007 in both species recorded and densities of freshwater, estuarine and marine species during the initial closed period was linked to increased salinity, lack of recruitment and reduced estuarine surface area. Increased wave action and high seas following Cyclone Gamede opened the mouth in March 2007 after a 57-month closed period, resulting in recruitment of marine species, predominantly P. indicus, followed by a decline in densities after reclosure in August 2007. Increases in marine and freshwater species were evident after Mfolozi flood connections during 2008, 2009 and 2010. Highest densities were recorded after breaching in 2007 and after flood connections, highlighting the importance of a marine link to maintain the recruitment of penaeid species.     

Chwaka Bay (Zanzibar,East Africa) as a nursery ground for penaeid prawns

The penaeid prawns are recruited in the sheltered sandy beaches and mangrove areas of Chwaka Bay (Zanzibar) at post-larval stage (7 mm). Year round incursions with a maximum during the warmer months of December to March were observed. February to March is considered as the peak recruitment period. Out of six species of penaeids represented in the area, Penaeus latisulcatus (75%) and Penaeus indicus (15%) were dominant. The recruitment pattern indicated greatest incursions of post-larvae with the flood spring tides of the night when the tidal flow is strong.The juvenile population of P. latisulcatus is distributed in those intertidal sandflats with a rich growth of seagrass and P. indicus in the muddy areas of mangrove forests. P. indicus showed affinity for euryhaline conditions, whereas P. latisulcatus showed no preference for lower salinity. Provision of food and shelter are considered as important factors for their nursery dependence. P. latisulcatus attained a size of 60–70 mm in five to six months and P. indicus 110–120 mm in six to eight months during their nursery phase. These juvenile penaeids were found to be omnivorous, feeding on animal products, plant material and detritus.The maturing P. latisulcatus emigrate back to the sea when they are about 65 mm, whereas P. indicus move out at about 120 mm. A positive correlation between post-larval recruitment and juvenile abundance was observed.     

Prawn community structure in the subtropical Mfolozi–Msunduzi estuarine system,KwaZulu-Natal,South Africa

The life cycles of many marine and freshwater prawn species are closely associated with estuarine habitat. Here, the prawn community of the Mfolozi–Msunduzi estuarine system is described and the system's potential as an alternative nursery for prawns during prolonged closure of the adjacent St Lucia estuarine system is examined. Sampling was conducted at five localities in summer and winter in 2007–2012. The Mfolozi–Msunduzi Estuary is a river-dominated system with mouth condition, sediment grain size and turbidity being the most important factors affecting the structure of its prawn community. Twelve prawn species were recorded, including five marine penaeid and five freshwater palaemonid species. The prawn community was numerically dominated by freshwater Macrobrachium equidens and two penaeids, Fenneropenaeus indicus and Metapenaeus monoceros. Significant differences were recorded between summer and winter samples, with freshwater species dominating the catch in summer, while penaeid species were dominant in winter. The marked seasonal change in the prawn community was related to strong river flows during summer creating low-salinity conditions, unsuitable for penaeid postlarval development, throughout most of the system. Compared to St Lucia, the Mfolozi–Msunduzi Estuary can be regarded as a poor alternative nursery area for penaeids during the summer peak postlarval recruitment period.     

Allopatric distribution of juvenile red-legged banana prawns (Penaeus indicus H. Milne Edwards, 1837) and juvenile white banana prawns (Penaeus merguiensis De Man, 1888), and inferred extensive migration, in the Joseph Bonaparte Gulf, northwest Australia

During October to December 1997, we trawled estuarine habitats in the Joseph Bonaparte Gulf (JBG) to determine the distribution of juvenile red-legged banana prawns, Penaeus indicus (H. Milne Edwards, 1837) and white banana prawns, Penaeus merguiensis (de Man, 1888). We made 229 beam-trawls at 185 sites, mostly over a 100-m path (3-min duration). A Global Positioning System (GPS) receiver was used to verify our location. During October to December 1998, we intensively resampled three of the rivers that were sampled in 1997 to confirm the gulf-wide distribution of P. indicus and P. merguiensis and to investigate the microhabitat use of P. indicus. We chose previously sampled and new sites in Forsyth Creek (eastern JBG), the Lyne River (Cambridge Gulf), and the Berkeley River (western JBG). We made 249 trawls at 21 sites, mostly over 100 m.Juvenile banana prawns were abundant in eastern JBG, Cambridge Gulf and western JBG. They were not abundant in southern JBG, although fewer trawls were made there, due to its inaccessibility. In eastern JBG and Cambridge Gulf, over 96% and 73% (respectively) of juvenile banana prawns were P. indicus and they were more abundant there than in the western JBG. Conversely, in the western JBG over 93% of the juvenile banana prawns were P. merguiensis and they were more abundant than in the eastern JBG and Cambridge Gulf. The Lyne River in the northwestern Cambridge Gulf seems to be the transition zone; both P. indicus and P. merguiensis are equally abundant.P. indicus are most abundant on the mangrove-lined muddy banks of waterways within mangrove forests, similar habitats to P. merguiensis. Within these habitats, they were most abundant in gutters and small creeks, rather than rivers and large creeks. Few P. indicus or P. merguiensis were caught in 100 m² trawls undertaken midriver (on the channel bottom and on emergent banks), although these habitats may be only 100 m from the mangrove-lined habitats. In all creek and river habitats, both species are most catchable at low tide (irrespective of daylight or darkness) when they move out of the mangrove forests and accumulate in the remnant water bodies.The offshore fishery for P. indicus is in northwestern JBG in waters 50-80 m deep, about 300 and 200 km, respectively, from where juveniles are abundant in their extensive inshore habitats in east JBG and in Cambridge Gulf, demonstrating a geographical separation of the juvenile and adult phases. Postlarval P. indicus, spawned offshore, must use tides and currents to travel south and east to reach nursery habitats. Emigrant subadults must migrate north and west, across relatively shallow inshore sand substrates (30-40 m deep) to reach their offshore habitats.     

Impact of a flood event on the zooplankton of an estuarine lake

Shallow coastal lakes are prone to large fluctuations in physico-chemical variables such as salinity and turbidity. This is now escalating in response to global change. A flood event in March 2014 resulted in a silt plume spreading through part of Lake St Lucia (South Africa). To determine the impact of this event on zooplankton, the Narrows region of St Lucia was sampled on a monthly basis from March to September 2014. For comparative purposes, data from samples collected prior to the flood event were included in the analyses. Analysis of similarity (ANOSIM) revealed dissimilarities in zooplankton community structure among the sampling occasions. The March 2014–May 2014 period was characterized by the highest abundance of freshwater species. Conversely, the abundance of the resident St Lucia copepods Acartiella natalensis and Oithona brevicornis was lowest during this time, and highest in September 2014. The other dominant copepod Pseudodiaptomus stuhlmanni prevailed in March 2014, but declined markedly in April. As of September 2014, P. stuhlmanni had yet to regain its pre-flood densities. The BIOENV procedure, which relates biological and environmental data, revealed that turbidity, salinity and dissolved oxygen were responsible for the observed changes in zooplankton community structure during the study period. Careful management of turbidity and salinity is stressed, as both factors are major drivers of the biota of St Lucia and similar systems worldwide.     

Endogenous swimming rhythms underlying secondary dispersal of early juvenile blue crabs, Callinectes sapidus

Blue crab, Callinectes sapidus Rathbun, megalopae settle in seagrass or other complex submerged aquatic habitats in estuaries, where they metamorphose to the first juvenile (J1) crab stage. Within tidal areas, early juveniles (J1-2) leave such nursery areas by undergoing secondary dispersal during nocturnal flood tides. The present study determined whether J1-2 blue crabs have a biological rhythm in vertical swimming activity that contributes to secondary dispersal. Endogenous rhythms in vertical swimming were determined for (1) J1-2 crabs collected from two estuaries with semi-diurnal tides, (2) J1 crabs that metamorphosed from the megalopal stage in the laboratory the day after collection, and (3) premolt megalopae that metamorphosed to J1 crabs under constant conditions during the experiment. In all cases, a circadian rhythm was present in which crabs swam vertically during the time of night in the field. The time of peak vertical swimming did not correspond to the time of flood tide at the collection sites, but did consistently occur at night, with a mean around midnight. While responses to environmental factors probably control the onset and end of vertical swimming by early juvenile blue crabs during flood tides in tidal areas, a circadian rhythm underlies secondary dispersal at night.     

2. NOTES ON THE STATUS OF SOME PALAEARC-TIC MIGRANTS IN NORTHERN RHODESIA

Whitfield, A. K. &; Cyrus, D. P. 1978. Feeding succession and zonation of aquatic birds at False Bay, Lake St Lucia. Ostrich 49:8-15.

Seasonal fluctuations in water levels at Lake St Lucia resulted in the inundation of depressions adjacent to the lake. Backwaters formed and were immediately colonized by aquatic invertebrates and fishes. Piscivorous birds dominated the early stages of the backwater feeding aggregations but with the decrease in water levels due to evaporation, crustacean and mollusc feeding birds became abundant. Both piscivorous and invertebrate-feeding wading birds formed a series of overlapping zones parallel to the shoreline.     

Larval and juvenile fishes occurring with flood tides on an intertidal mudflat in the Tama River estuary, central Japan

Intertidal movements of fish larvae and juveniles on a mudflat in the Tama River estuary, central Japan, were investigated by comparing the abundance and sizes of fishes caught in the intertidal zone during flood tides with those in the subtidal zone during low tides. A total of 28465 individuals, belonging to 9 families and 20 species, were collected by small purse seine. Among the abundant species, planktonic larvae and juveniles of gobiids and Konosirus punctatus were more abundant in the intertidal zone at flood tide than the subtidal zone at low tide. Similar occurrence patterns were found in juvenile Plecoglossus altivelis altivelis and Lateolabrax japonicus, having fully developed swimming abilities. In contrast to these species, much higher abundances of epibenthic juveniles of 2 gobiids (Acanthogobius flavimanus and Gymnogobius macrognathos) were found in the subtidal zone at low tide, although they also utilized the intertidal zone at flood tide.     

In situ feeding rates and grazing impact of Mesopodopsis africana O. Tattersall in the St Lucia Estuary, South Africa

Mesopodopsis africana is an important mysid in southern African coastal zooplankton and a key species in the St Lucia estuarine lake, which is currently undergoing severe desiccation owing to freshwater deprivation. M. africana populations through much of the system are consequently under severe environmental stress. This study investigates the grazing dynamics of this mysid species, in relation to autotrophic food availability and other environmental constraints in two contrasting areas of the St Lucia Estuary, Charters Creek, heavily affected by the desiccation process and the Mouth, virtually under unchanged conditions. Gut evacuation experiments were conducted once each during the day and the night. Evacuation rates were consistently higher during the night, ranging from 0.27 to 0.33 h^-1 at Charters Creek and from 1.13 to 1.24 h^-1 at the Mouth. Ingestion rates were, therefore, higher at the Mouth resulting in population grazing impacts of 2.5% of the total microalgal biomass, while the grazing impact at Charters Creek was only 0.5%. The spatial variation in ingestion rates could be attributed to seasonal differences in gut evacuation rates, differences in the mean size of mysids used, or the physicochemical conditions present at the two stations. It is suggested that mysid populations at Charters Creek are predominantly driven by bottom-up forces, initiated by the harsh environmental conditions. Despite the lower ingestion rates exhibited at Charters Creek, results indicate that these mysids are capable of meeting all their energetic requirements from a microalgal diet alone, although they may also utilise a heterotrophic diet.     

Tidal and diurnal variations in larval fish abundance in an estuarine inlet in Ariake Bay,Japan: implication for selective tidal stream transport

We have conducted a preliminary study of tidal and diurnal variations in the distribution of dominant larval and juvenile fishes in the Chikugo River inlet (Ariake Bay, Kyushu, Japan) to determine whether selective tidal stream transport (STST) occurs. Larval and juvenile fish were collected from the mesohaline zone of the Chikugo River inlet during spring 2002. Temperature, salinity, depth, and current velocity were measured. Larval and juvenile abundance were compared among four tidal conditions, flooding tide, high tide, ebbing tide, and low tide, and between day and night. A total of 12 families, 15 species, and 5,577 individuals were collected. Temperature did not vary significantly with tidal conditions whereas salinity, depth, and current velocity varied significantly. Salinity also was correlated significantly and positively with depth. The abundance of most of the fishes was correlated positively and significantly with salinity and depth. Lateolabrax japonicus, Trachidermus fasciatus, Acanthogobius hasta, and other gobiid larvae (Gobiidae spp.) were significantly more abundant during high tide; in contrast, Coilia nasus and Neosalanx reganius were most abundant during low tide. The abundance of most of the fishes was higher during high tides at night than during the day, indicating the existence of STST, which may be strategically associated with ascending progress to upstream nursery areas.     

Assimilation efficiency of zinc and cadmium in the decapod crustacean Penaeus indicus

The assimilation of trace metals from food can be the main route of metal uptake into aquatic invertebrates. The assimilation efficiencies of zinc and cadmium from muscle tissue of a cephalopod mollusc (40% to 70% Zn, 64% to 83% Cd) and from a macrophytic alga (50% to 69% Zn, 39% to 50% Cd) were measured in juvenile penaeid prawns Penaeus indicus. Assimilated Zn and Cd were retained mostly in the hepatopancreas of the prawns, some metal being excreted (Zn k_e 0.10 and 0.11 and Cd k_e 0.004 and 0.009, from cephalopod muscle and alga, respectively). There were no significant differences between k_es (efflux rate constants) of one metal from either diet. Given the high trace metal assimilation efficiencies measured here, it is highly probable that metal assimilation from food plays an important role in Zn and Cd accumulation in the body of P. indicus, particularly in estuarine stages of the life cycle, for estuaries are particularly prone to metal pollution and are likely to offer the prawns abundant metal-rich diets such as detrital material derived from local macrophytes.     

Foraging strategy of the mudskipper Periophthalmus sobrinus Eggert in a Kenyan mangrove

The feeding behaviour of Periophthalmus sobrinus was studied in a population living along the channels of the Tana river delta. In November 1993 mudskippers were sampled every 2 tidal h for two consecutive spring and neap tides both during the day and night. The fishes were sexed and their stomach contents analysed. The results indicated that the mudskipper has a carnivorous diet and forages during both day and night although feeding greatly decreases after dark. Quantitative and qualitative differences in diet were found during the day and night of the two synodic and tidal phases. These differences were also demonstrable in the two sexes. The study showed that females of P. sobrinus are more selective than males and that differences between the two sexes may be related to reproductive condition. Furthermore, the mudskippers showed a tidal rhythm in feeding behaviour with peaks of activity around low tide (LT) on spring tides and around high tide (HT) on neap tides. Foraging activity seems to be influenced by the different spatial and temporal distributions of the mudskipper's prey caused by synodic, tidal and diel changes in the environment.     

The distribution and abundance of planktonic penaeid larvae in Tudor creek,Mombasa, Kenya

The temporal distribution and abundance of penaeid prawn larvae in surface waters of Tudor creek were studied. Only late mysis and early post mysis stages of development were encountered in the creek waters with a descending gradient from the mouth station (station 1) to the innermost station 5. Larval encounter incidence (% incidence) was highest in plankton tows taken during night spring tides (59%) followed by night neap tides (45.4%), day spring tides (34.9%) and day neap tides (21.4%). This encounter incidence was more skewed towards the seaward stations (1 and 2) during the neap tides than during spring tides. Likewise, larval density (mean catch/standard 5 min. tow) decreased less steeply up the creek during spring tides than it did during neap tides. Higher catch rates were in general obtained in night samples than in day samples at each station and in each neap/spring phase. The diel cycle had a greater effect on both incidence and abundance of larvae than did the tidal cycle.Intensified larval incursion into the creek was observed between March and June.     

Body temperature and basking behaviour of Nile crocodiles (Crocodylus niloticus) during winter

The ability to thermoregulate in reptilians is often through behavioural modification. We investigated body temperature (T_b) patterns during winter in the amphibious Nile crocodile (Crocodylus niloticus) and its relationship to basking behaviour at the St. Lucia Crocodile Centre, St. Lucia, South Africa. It was found that crocodiles had no daily plateaus in T_b but rather continuous oscillations in T_b within a range of mean minimum T_b 18.8–19.6 °C to mean maximum T_b 26.9–29.2 °C. Crocodile T_b increased during the day, usually after 10:00 irrespective of body size. Behavioural data showed that the crocodiles usually left the water to bask around 10:00. It is suggested that basking behaviour is important for elevating T_b rather than attaining a preferred T_b. The increased T_b may allow them to perform optimally when they return to water. The basking occurrence has management implications as it suggests that the best time to conduct aerial censuses of the St. Lucia crocodiles is during winter after 10:00 when most of the individuals are basking and hence most easily seen.     

Thermal tolerance of Litopenaeus vannamei (Crustacea: Penaeidae) acclimated to four temperatures

Critical thermal minima (CTMin) and maxima (CTMax) values were determined for the Pacific white shrimp Litopenaeus vannamei post-larvae and juveniles at four different acclimation temperatures (15, 20, 25, and 30 °C). The CTMin of shrimp at these acclimation temperatures were 7.82, 8.95, 9.80, and 10.96 °C for post-larvae and 7.50, 8.20, 10.20, and 10.80 °C for juveniles, respectively, at 1 °C h⁻¹ cooling rate. The CTMax values were 35.65, 38.13, 39.91, and 42.00 °C for post-larvae and 35.94, 38.65, 40.30, and 42.20 °C for juveniles at the respective acclimation temperatures. Both acclimation temperature and size of the shrimp affected CTMin values of L. vannamei (P<0.01). Overall, juveniles displayed significantly lower CTMin values than the post-larvae (P<0.0001). However, the CTMax response by post-larvae and juveniles were not significantly different from each other and no interaction was determined between the acclimation temperature and development stage (P>0.01). The area of the thermal tolerance polygon over four acclimation temperatures (15, 20, 25, and 30 °C) for the post-larvae of L. vannamei was calculated to be 434.94 °C². The acclimation response ratio (ARR) values were high ranging from 0.35 to 0.44 for both post-larvae and juveniles. L. vannamei appears to be more sensitive to low temperatures than other penaeid species and its cold tolerance zone ranged from 7.5 to 11 °C. In successful aquaculture temperature must never fall below 12 °C to prevent mortalities. Upper thermal tolerance is less of a problem as in most subtropical regions maximum water temperature rarely exceeds 34 °C, but care should be given if shallow ponds with low water renewal rate are being used.     

Molecular phylogenetic analysis of white prawns species and the existence of two clades in Penaeus merguiensis

Closely related species, Penaeus merguiensis and Penaeus silasi from Thai waters, were genetically examined using variation observed in 558 base pairs (bp) of sequence from cytochrome oxidase subunit I (COI) gene of mtDNA. The sequence divergences of COI between P. merguiensis and other Penaeus species were 5.76-6.15% (P. silasi), 13.17-13.97% (Penaeus indicus), 16.43% (Penaeus vannamei), 16.63% (Penaeus monodon), and 18.37% (Penaeus japonicus). From the alignment reported that there were four clades on phylogenetic tree, the distinction of the two monophyletic clades was referred as P. merguiensis, one monophyletic clade within P. silasi and P. indicus. These results point toward the possibility of P. merguiensis being a complex of two cryptic species or a single species with strong phylogeographic subdivision.     

Diel and tidal changes in the distribution and feeding habits of Japanese temperate bass Lateolabrax japonicus juveniles in the surf zone of Ariake Bay

Japanese temperate bass Lateolabrax japonicus juveniles recruit to the surf zone and grow by feeding on commonly occurring coastal copepods. However, little is known about diel and tidal patterns in their migration and feeding habits. We sampled wild juveniles during the neap and spring tides, over periods of 24 h, with small seine nets in the sand flat of the eastern part of Ariake Bay, Kyushu, western Japan. In both the neap and spring tides, abundance of juveniles significantly increased during the daytime, being highest around the time of high tide. The relative gut fullness indices of juveniles drastically increased in the morning (0700–0900) and during the flood tide in the daytime, while major prey copepod (Paracalanus spp.) density in the ambient water was relatively constant. We summarized that L. japonicus juveniles would migrate to the surf zone after sunrise to feed on copepods, and then emigrate from the surf zone after sunset. The migratory behavior of L. japonicus juveniles would be influenced by light (daytime) and feeding activity influenced by both light (morning) and tidal condition (flood tide). The intertidal region of the tidal flat was recognized to be one of the important habitats for L. japonicus during their early life history.     

Patterns of temporal occurrence of brachyuran crab larvae at Saco mangrove creek, Inhaca Island (South Mozambique): implications for flux and recruitment

Temporal patterns of larval occurrence of brachyuran taxa weredescribed from Saco mangrove creek, Mozambique, based on planktonsamples. Brachyuran larvae were collected hourly in four 24h cycles during neap and spring tidal periods at a fixed station,in November 1997 and February 1998. Results indicate a semilunarcycle of larval release activity for most species. Newly-hatchedstages of sesarmids showed a peak occurrence in post-crepuscularebbing tides, Macrophthalmus spp. and Uca spp. showed highestdensities during ebbing tides at night. Dotilla fenestrata andPinnotheridae, in spite of showing a semilunar pattern, didnot display significant differences between day and night. Mostmangrove taxa showed thus a clear larval exportation behaviourduring ebb tides, with little return during the following floods,with varying degrees of dependence on the diel period. Megalopaewere restricted to the night period, and highest values wereobtained during spring tidal periods. However, maximum valuesof megalopae were obtained both during flood and ebb periods,contrary to most previous studies. This is interpreted as abi-directional transport of non-competent megalopae into andfrom the mangrove area.     

Genetics and shell morphometrics of assimineids (Mollusca,Caenogastropoda, Truncatelloidea) in the St Lucia Estuary,South Africa

The Assimineidae are a family of amphibious microgastropods that can be mostly found in estuaries and mangroves in South Africa. These snails often occur in great numbers and are ecologically important to the St Lucia Estuary, which forms a crucial part of the iSimangaliso Wetland Park, a UNESCO World Heritage Site. Genetic and shell morphometric analyses were conducted on individuals collected from nine localities distributed from the northern lake regions to the southern lake and the mouth of the St Lucia estuarine lake. Mitochondrial (COI) and nuclear (28S) DNA was used to construct Bayesian Inference, Neighbour-joining, Maximum Parsimony and Maximum Likelihood trees. Principal Component Analysis and Cluster Analysis were performed on standard shell parameter data. Results indicate that two different taxa are present in St Lucia. The taxon comprising individuals from the South Lake and St Lucia Estuary Mouth is identified as Assiminea cf. capensis Bartsch, in accordance with the latest taxonomic consensus. The taxon comprising assimineid individuals from False Bay, North Lake and South Lake, is here tentatively named “Assiminea” aff. capensis (Sowerby). These two taxa exhibit patterns of spatial overlap that appear to vary depending on environmental parameters, particularly salinity. The need to resolve the complex taxonomy of assimineids is highlighted.     

Biodiversity census of Lake St Lucia,iSimangaliso Wetland Park (South Africa): Gastropod molluscs

The recent dry phase experienced by the St Lucia estuarine system has led to unprecedented desiccation and hypersaline conditions through most of its surface area. This has changed only recently, at the end of 2011, with the onset of a new wet phase that has already caused a major shift to oligo- and mesohaline conditions. The estuary mouth, however, remains closed to the ocean, making the weak connection recently established between the St Lucia and the Mfolozi estuaries the only conveyance for marine recruitment. As a result, only 10 indigenous and two alien aquatic gastropod species are currently found living in the St Lucia estuarine lake. This is out of a total of 37 species recorded within the system since the earliest survey undertaken in 1924, half of which have not been reported in the literature before. The tick shell, Nassarius kraussianus, which was consistently found in large abundance prior to the recent dry phase, appears to have temporarily disappeared from the system, probably as a result of the extinction of Zostera marine grasses inside the lake. Population explosions of the bubble shell Haminoea natalensis, with its distinct egg masses, were recorded seasonally until 2009, but the species has subsequently not been observed again. A molecular DNA analysis of the various populations previously reported as belonging to the same assimineid species, variably referred to as Assiminea capensis, A. ovata, or A. bifasciata, has revealed that the St Lucia assemblage actually comprises two very distinct taxa, A. cf. capensis and a species provisionally referred to here as “A.” aff. capensis or simply Assimineidae sp. In the mangroves, the climbing whelk Cerithidea decollata is still found in numbers, while ellobiids such as Cassidula labrella, Melampus semiaratus and M. parvulus are present in low abundances and all previously recorded littorinids have disappeared. A number of alien freshwater species have colonized areas of the system that have remained under low salinity. These include the invasive thiarid Tarebia granifera, which can be found in concentrations exceeding 5000 ind.m^-2, the lymnaeid Pseudosuccinea columella and the physid Aplexa marmorata.